l

The State of Phtllptne Herpetolagl 19

THE STATE OF PHILIPPINE HERPETOLOGY AND THE CHALLENGES FOR THE NEXT DECADE Daring tbe tatne perirtd. tbere bat been let.r actiuity in ecological research and conteruatittu. ancl little or no actiuity in disciplines tach at behaaior, nicroeao/ution, reproductiue Reru M. BnowN, Anvnv C. Dtrslros, aNo ANcn C. biol- lgy, 0r popalation biologl'. Ix thit paper we reaiew a nodel Ar-cnt-e fetu studies and point out where others are hadly needed. Auailable biogeographic analyset, combined with new, First, there is a great need for more new basic re- unpa[tlished data, dent.on.rtrclte thdt the distributions of anphib- searc h focused on b iodive rs i ry conservat io tt, includittg system- iant and reptiles in tbe Pbilippines haue been ttrongly influ- atics, ecology, behavior, and current patterns of distibution enced by the nid- to late-P /ei.ttlcene format ion of seuera/ aggre- and ab u ndanc e. W itho ttt s uc h fundantenta I info r nat io n, c o n- gate island complexes at well at by clinatic gradients attociated servatton planning will be incornplete at best. with eleaation and a,ttbrlp0geTtic distarbances (priraarily cle- Heaney et al., 1999:315. forestation). Each Plei-rtocene aggrega.te island complex it a major center of biologica/ diaersity, and within tbese najor (and seu- The information needed to make sense of Asian eral other rninor) Iand nass amalgamatilnt, there exist numer- herpetology is not lurking in tlrc Literature; it is still out tlrcre in ous tub-centers of endernism and diuersitl centered on itolated tlrc rice padtlies and in the vanishing patches of nontane for- ntountains or mountain ranget. Ampbibians and reptiles rnay est. represent particalarly appropriate model organismt for the study Crombie. 1992:593 of tbese lerser centert of biologica/ organization due to tbeir ten- dency towardt finer-scale differentiation and isolation on tingle nzrntane "itlandt" and mountain ranget. Seueral recent studiet ABSTRACT haue began the process of integrating phylogenetic data, tpecies distribution data, and ttudies of tbe pr0ce.rr of speciation on uniqae nrntane habitatt, ltat rnany ,nrre dre needed. ln par* ,Tht herpetological faana (amphibiau and repti/et) of the Philip ticalar, tbe of nolecular s)ttlematicJ ttandt ()ut.tJ an im- I pines h extrenely rich in total species numbert, taxononic diuer- field mextely powerful tet of tool.t tbat ba.r yet to be ,apped by conter- sity, and percent endenism----especially when considered as afunction of aati0n biologists in the Philippines. aaailable land area. Tbe last 1 0 years of berpetological retearch in The last clecade hat .seen .reueral attemptt t0 ar,ieJr the the Philippines haue Jeen a dramatic inuease in interest in tax- clnJeraat;0n and pre- rnrrny, biogeq%phy, phylogenetic systernatict, conseruation, and !tatilr of nanl of the Philipltines' unique sumably tltreatened arnpbibians and reptiletJbzse effort.s baue biodiuenitT of Philippine tpeciet, especially atnphibians. ln tbe been bdrnpered by a general lack of knowledge, a paacity of batic last decade, oaer 50 prcuioutly uarecognized species baae been bateliae saraey data, a lack of integration, public disinterest, identified. Despite the publication of a recent field gaide to the bareaacratic obttacles to rctearcb, and b1, linitationt in retourceJ. anphibians of the Philippines, auailable speciet tumuaries antl The namber 0ne ca,tre of anphibian and reptile population de- diagnostic Aeys are cuffently oat of date because progress has clines clearly is catastrophic habitat deJtructi0n due to thc ac- been so rapid. Reuisions of these urrkJ are needed but tnust await tiuities of hamans. the completion of seueral conprehensiae taxonomic inuestigations cutrently in progrett. In general, ampbibiant (especially ranid frogs) haue receiued nore dttenti0n tban reptilcs.

^9illiman,Iournal Vol. 42 No. I 2001 Silliman "Iournal Yol.42 No. I 2001 I ll/3 JtuLe uJ I tLLLLPLtLc L t't ycLvw6J - ^ 20 Brown, Diesmos E Ahaln

Introduction Compositionofthelastl0years'publishedliterature

we considered only published papers Situated at the interface between the Oriental and For this review writing, accepted or in press) Australian faunal zones is the largely oceanic island (or ones that were, at the time of M'S' and/or Ph'D' nation of the Philippines. The Philippine islands are and unpublished undergraduate honors, data (theses and a few pa- home to a spectacular and diverse set of amphibian and theses. We mention unpublished in some cases but we can in- reptile radiations that have captured the attention and pers in review or preparation) private papers' or other imagination of diversity specialists and biogeographers contracte-d reports, "t.ra" not been or will not be peer re- since the first accounts of Philippine herpetological di- pseudopublications that have (see Literature cited section) versity appeared in the scientific literature (e.g', viewed. wehave compiled 109 Philippine herpetology from be- Boulenger, 1882, 1894, T920; Peters, 1863; Boettger, scientific publications on 2001 (Fig. 1). The annual publica- 1893; Taylor, 1915, 1918a, 1919,1920a,1920b, 1921, tween the years 1990 and 1922a, 1922b; Taylor and Noble, 1924; Noble, 1931 ; tionratehasremainedrelativelystable,withnotableexcep- Schmidt, 1935). The career of Edward H. Taylor in the tions(i.e.,inlgg5numerousarticleswerepublishedonrep- were published on amphibians in 1920's (Taylor, 1975) brought the Philippines to the fore- tiles while many articles front of global appreciation of amphibian and reptile 1999and2000).Thecompositionofthelastdecade'spub- research in sys- diversity as one of the world's major centers of lished record was markedly skewed towards and species diversity (Fig' herpetological diversity and endemism ' Later taxonomic tematics, taxonomy, biogeography, studies consisted of and biogeographic summaries (Inger, 1954, 1999; 2). The vast majority oi the remaining population biology and community stud- Leviton, 1963; Alcala, 1986; Brown and Alcala, 1970a, ecological (includes and only a very small fraction I978, 1980, 1994;Allison, 1996;Brown, 1997; Alcala ies; aid conservation studies (e.g', information on Quaternary and Brown, 1998) further promoted the recognition of addressed other subjects Reis, 1999; Reis and Garong' the importance of Phllippine herpetological diversity and herpetofaunal communities; articles that, in part' addressed stressed the unique nature, evolutionary history, and 2001) or were popular of amphibians and rep- remarkable diversity of Philippine amphibians and rep- herpetological topiis or biodiveisity Diesmos, 2000, 2001; Brown and tiles (see also Noble, 1931; Euellman and Trueb , 1994)' tites (geaney et al., 2000; aL The last l0 years in Philippineherpetelogieakesearch Alcala, 2000; Brownct J002)' have seen an increase in interest in a diverse range ofstudies studies and species diversity in set against the backdrop of an emerging period of unprec- History of herpetological edented taxonomic rediscovery, concern for conservation, and the PhiliPPines an increase in appreciation for biodiversity. The purpose of published papers on Philippine herpetology this paper is to review and analyze the past decade's progrgss' The first Boettger, Boulenger, Gtinther' Mertens' to consider its significance within the context of the history included the works of among others (see Inger' of Philippine herpetology, and to identify prospects and Peters, Weigmann, and Stejneger' Brown and Diesmos' this goals for future research and conservation. 1954;Bayleis and Adragna,1997

Yol' 42No. 1 2001 Silliman,/ournal Yol.42 No. 1 2001 ,silliman,/ournal volume). This "age of discovery,' philippine in herpetology The work of A. Alcala and W. Brown later sct thc marked the first exposure philippine of the outside worlcl to stage for present studies that continue in collaboration with herpetological diversity, and the papers that resulted were al- A. Diesmos and R. Brown. Currently, we recognize a total of most entirely descriptive in nature. The first worker to con- 101 species (78, or 77o/o, endemic) of Philippine amphibians centrate efforts on a comprehensive philippine review of (Fig. 3) and an approximate total of 258 (169 or 659,i, en- herpetofauna was Edward Harrison Taylor (1915-lOZS, see demic) species of Philippine reptiles (Fig. a), That estirnate Literature cited). In his numerous taxonomic works, Taylor will surely increase by 10-20% in the coming years as nu- recognized a total of 89 amphibians and approx imately 253 merous undescribed species are named in ongoing taxonomic reptiles. Later,Inger (l 954,1960a. 1960b; see also Hoogstral, reviews (R. Crornbie, pers. comm: Diesmos, Brown, and 1951) recognized 55 species of philippine Amphibia, reduc- Alcala, unpublished data). Surnmaries of taxa described in ing the species level diversity of philippine Amphibia by ap_ the last decade are presented in Tables 1 and 2. plication ofthe Polytypic Species Concept (see Brown ,199i; The vast majority of papers during the last l0 years Brown et al., 2000; Brown and Diesrnos, this volume). In the of progress in classification and recognition o1'Philiprpine mid-1950s Angel Alcala and walter Brown began a collabo- herpetological diversity have been species descriptions (c.g.. rative review of most major groups of (see lizards also Inger, Ota and Crombie, 1989 Lazell, 1992; Wynn and Ler,'iton. 1958, 1983; Musters, i983; Inger and Brown, l9g0) in the 1993; Alcala et al., 1998; Brown et al., 1995a,1999a,1999b; Philippines and during the course of their pub_ field work, Brown et al, 7997 c, 1999a, I 999b; Larua, I 999; Gaulk e,2002, lished numerous additional species descriptions (see Liiera- Diemos et al., in review), redescriptions of poorlyunderstood ture Cited: Alcala, I 955-l 986; Alcala and Brown, l9S5_I999; taxa (Ota et a1.,1993; Brown et a1.,1997; Brown et al., 1998), Brown and Alcala, 1955-1994; Brown et al., lgg1_lggg). or clarifications of species boundaries (Ota et al., 1989; Brorvn During the sameperiod, Alan Leviton systematically et al., 1998; Brown et al, 2000a,2000b, 2000c, 2001: reviewed the contents ofmost philippine snake genera in his Gumprecht, 2001). Additionally, several important papers contributions to a review of philippine snates series (Leviton, have taken the form of rnore comprehensive reviews of gen- 1955-1983; see also Leviton and Brown, l95g; Inger and era or species groups (Gaulke 1992a: Dubois, 1992; Ota and Marx, 1965; Inger and Leviton Gyi, 1970; ,1966; McDowel, Ross, 1994; Inger, 1996; Bayless and Adragna, 1997; Fritz et 1974;Malnate and Underwood, l ggg). Alcala (19g6; see also a1.,1997; Brown et al., 1997a,1997b,1999b; Brown et al.. Rabor, 1981) summarized some ofthis taxonomic work, rec_ 2000a,2000c; Brown and Diesmos, this volume; Brown and og-nizing 66 arnphibian +nd 205 reptile species (see also GUtman, in press; McGuire and Alcala, 2000; Dubols ancl Afuang, 1995; Gonzales, 1995; DENR and UNEp, lggT). Ohler, 2000; Veith et al., 2000; Helfenberger, 2001). All of Progress was made towards a synthesis of spe.cies di_ these studies have greatly increased recognized species cli- versity by the unpublished works of R. I. Cromb'ie Qcers. versity in the Philippines. comm.). Crombie's bibliography and annotated check_ In amphibians, the greatest areas of activity have been list have served as the backbone of many working spe_ in ranid frogs. For example, in the Rana signata and Rctna cies lists used by researchers in the philippiner in it everetti species groups, diversity has increased frorn two to past decade. " twelve species (Brown et al, 2000a; Brown and Diesmos. this

Silliman "/ournal Yol.42 No. 1 2001 ,Silliman "/ournal Yol.42 No. 1 2001 '25 lhE State ol Fhilipine Herpetolngy

volume; Brown and Guttman, in press) and plafymantine ranid surnably closely related to the Philippine endemic Varanus frog diversity has increased from seven (lnger 1954) to more olivaceus,has captured the attention of herpetologists around than 25 species (Alcala and Brown, 1998, 1999). We now the world. These studies indicate that an enorrnous amount know that the species diversity of Philippine flying lizards of descriptive taxonomic work has yet to be conducted in the (genus Draco, l0-I2 species; McGuire and Alcala, 2000) is Philippines before we can adequately assert that the coun- closerto original estimates of Taylor (l922a,who recognized try's amphibian and reptilian speoies diversity is reasonably 11 species) than it is to later estimates of Inger (19g3), who well known. recognized three species (see also Musters, 19g3). In total, The types of data utilized by amphibian and reptilian over 50 previously unrecognized species have been identi- taxonomists working in the Philippines have changed in some fied in the past decade. Thirty-two of these have been for- cases but have remained the same in many others. Although mally named or resurrected from the synonymies of wide- taxonomists are now distinguishing between species with spread polytypic species complexes (14 reptiles and l8 frogs). DNA sequence divergence data (McGuire and Kiew,200l; At present, more than 15 endemic philippine frog species Brown et al., unpublished data), phylogenetic evidence such await description (Diesmos, Brown, and Alcala, unpublished as a species' position in evolutionary trees (McGuire and data), and we suspect that many more await discovery. Alcala, 2000; Brown and Guttman, in press), fixed allozyme Some recent discoveries have been truly spectacular. differences (Brown, 1997; Brown and Guttman, in press), A new, very distinctive, endemic philippine genus ecological differences (Brown et al., 2000a, 2000c) and (Parvoscincus) of scincid lizards was discovered in the last behavioral differences (especially variation in acoustical ad- decade (Ferner et al., 1997), and further generic subdivision vertisement si gnals of male frogs ; Brown et al., I 997 c, I 999 a, of one group of ranid frogs currently is underway (Brown et 1999b; Brown and Guttrnan, in press), the majority of recent al., unpublished data). Recognition of philippine taxonomic papers have used morphological data in the tbrm herpetological diversity has not simply been aprocess ofsplit- of character differences and comparisons of morphornetric ting closely-related species; in fact, higherlevels oftaxonomic measurements or ratios of body proportions (Brown et al., diversity are poorly understood in several key areas. The 1997a, I997b, 1997c, 1999a; Brown et al., 1995a, 1995b, phylogenetic affrnities e o s of H e rny s (: " G e o my d a' ) I ey t e ns i s 1999a, 1999b, 2000a, 2001 ). and H. spinosa are unclear; generic revision of these taxa may be required with on-going systematic studies (see lgylAf, Review of biogeographic studies of Philippine amphib- -TyzO;F, Alcala, 1986; Timmernan and Auth, 19gg; Buskirk, ians and reptiles 1989; Iverson, 1992; Das, 1996a; Shaffer et al., 1997; Gonzales et al., 1997; McCord et al., 2000). A separate ge- The first attempt at a biogeographic summarization nus, Coelognathus, has been resurrected to accommodate of Philippine herpetofauna was Taylor's (1928) chapter in Indo-Malayan ratsnakes (previously of the genus Elaphe; Dickerson's Distribution of Life in the Philippines. Taylor [Leviton, 197 9J),including four philippine taxa (Helfenberger, (1928) summarized the known species diversity at the tirne, 2001). Finally, the discovery of a spectacular new species of plotted the distribution of the genera throughout the archi- frugivorous monitor lizard (Gaulke and Curio, 2001), pre- pelago, and commented on possible dispersal routes. He also

,Silliman ulournal Yol.42 No. 2001 I ^Silliman uloumal Vol.42 No. I 2001 r 26 Bro*n, Diesmos E Ahala Tfu State of PhiLipine Herpetobgy 27 recognized the distinction between land-bridge (e.9., Palawan Camiguin or Tablas; Brown and Alcala 1967.1974, 1978) Aggregate Island Complex) and oceanic portions (the remain- the explicit geological basis for the processes that may have der) of the Philippines, although his distinction was inferred led to these patterns of species endemism had not been em- from distributional data from the fauna and not explicitly from phasized. However, although Inger (1954), Leviton (1963), a knowledge of channel depths or geological reconstructions. and Brown and Alcala (1967, 1970a, 1986) acknowledged Taylor also noted the presence of several Sunda Shelf taxa in channel depths as potential barriers to dispersal (deeper chan- Palawan herpetofauna and the distribution of the more spec- nels indicative of a reduced chance of landbridges having tacular Philippine radiations (lizards of the genus existed in the past), the underlying framework for recogni- Brachymeles, frogs of the genus Platyntantis, and snakes of tion of all deep water islands as unique centers of biological the genera Oxyrhabdir.tm, Cyclocorus, and Hologerrhunz) con- endemism was not widely recognized until Heaney (1985, fined to the oceanic portions of the Philippines. 1986) traced the underwat-er 120 m bathymetric contours Later biogeographic sumrnaries included papers by throughout the Philippines (Fig. 5). This exercise explicitly Inger (1954,1999) on amphibians, Leviton's (1963) paper on illustrated Pleistocene sea shores at the end of last glacial snakes, Brown and Alcala's (1978) comments on gekkonids episode (22-12,000 years before present) and the formation and their summary of the biogeography of the archipelago's of enlarged aggtegate island complexes by exposure of land herpetofauna (Brown and Alcala, 1970a). Brown (1997), positive connections between Philippine islands separated by Allison (1996), and Inger (1999) have summaizedthese data less than 120 m (Fig. 5). The recognition of Pleistocene ag- in the larger context of SE Asia and the SW Pacific. Most of gregate island complexes is the appropriate framework for these studies take similar approaches, namely the discussion appreciation of Philippine biodiversity on all levels (Heaney of the zoogeographic relationships of the islands as indicated and Regalado, 1998), for it is the unique geological history by calculation of faunal similarities (see also Brown and of the islands that unites the evolutionary histories of all these Alcala, 1986 and Femer et al., 2001). All of these traditional islands' residents (review: Brown and Diesmos, this volume). summaries recognized most of the faunal subprovinces (five Understanding of mid- to late-Pleistocene geology is the key to seven distinct Pleistocene Aggregate Island Complexes) to appreciating the distribution of life in the Philippines ofHeaney (1985, 1986) as unique centers ofbiological ende- (Taylor, 1928; Inger ,1954; Leviton, 1963; Brown and Alcala, mism. Thus, Inger (1954), Leviton-(1963) and Brown and 7970;Heaqgy, 1985, 1986; see also Hall, 1996, 1998), and it Alcala (1970) all taxonomically recognized suites of endernic is the key to formulating effective conservation strategies taxa on Luzon as separate from those of Mindanao or the (Utzumrm, 1991; Oliver and Heaney, 1997 Heaney and Visayas (as embodied by the known heqpetofauna ofNegros; Regalado, 1998). Additionally, interpretation of Philippine see Ferner et al, 2001) but fell short of acknowledging the biodiversity in the context of Pleistocene geology is the best importance of the lesser studied deep water islands,of approach for formulating taxonomic and zoogeographic hy- Mindoro, Sibuyan, Siquijor, Tablas + Romblon, Burias, is- potheses (see below) for testing in a phylogenetic context lands of Batanes and the Babuyans, Camiguin, and Lubang. (Brown, 1997; Brown et al., 2000c; McGuire and Alcala, So, although endemic species were described from some of 2000; McGuire and Kiew, 20Ql; Brown and Guttrnan, in these islands (e.g., frogs and gecko endemics of Babuyans, press).

Silliman,Iournal Yol.42 No. 1 2001 ^Silliman,Ioumal Yo1.42 No. 1 2001 LA DTOWfu, litesmos Ct .I\LCAIA

studies of Philippine Finally, one last class of papers warrants con- Phylogenetic and phylogeographic sideration when reviewing Philippine biogeographi- amphibians and reptiles cal studies. These are faunal inventories, focused on singular sites or regions (i.e., Leviton, 1955; The last several years have seen the advent ofa new Brown (1997; Alcala, 1956, 195 8; Rabor and Alcala, T959; group of studies in Philippine herpetology. Alviola et al., 1998; Smith, 1993a, 1993b; Ubaldo, Brown and Guttman, in press) conducted the first phylogenetic Philippine amphibians, 1999; Reis and Garong, 2001), particular mountains analysis of an endernic radiation of or mountain ranges (Alcala and Brown, 1955; and Brown et al. (2000c) and McGuire and Kiew (2001) pub- Custodio, 1986; Alcala et al., 1995; Brown et al., lished the first phylogenetic analyses of SE Asian reptiles with 1996;2000b; Diesmos, 1998), small islands (Brown a significant proportion of their diversity represented in the and Alcala, 1963b, 1967, 1974; Ross and Lazell, Philippines. These three studies are significant in that they 1991; Ross and Gonzales, 1992; Gaulke, 1993, represent the first of their kind in Philippine herpetology and 1999; Gaulke and Altenbach, 1994; Gaulke, 1994a, also because they strongly support interpretations of 1995a, 1996,L999), and large islands (Gaulke, biogeographic patterns and routes of island colonization not 1994b, 2001a,2001b, 2001c; Sison et al., 7995; previously suggested by data from birds and mammals. For example, Brown (1997) found that the Philippine Rana sig- Denzer et al. 1999; Ferner et al, 2001; Gaulke ,. 2001a,2001b,2001c). One important new study (a nata complex was composed of two major clades of frogs first of its kind in Philippine herpetology) addressed (Fig. 6a), one centered on the eastern Philippine island arc biogeographical relationships of Palawan using new (Sulu-Mindanao-Leyte-Samar-Luzon) and one centered on the Brown, data on late Quaternary vertebrate communities, western island arc (Palawan-Buswanga-Mindoro; including amphibians and reptiles (Reis and 7997;Brown and Guttman, in press), and that the stream frogs Garong, 2001). Further faunal inventories are badly from Mindoro island were more closely related to those from needed to fill in gaps in distribution data left by Palawan and the Sunda Shelf than they were to the entire earlier biogeographic summaries that conspicuously remainder of the oceanic portion of the Philippines (contra missed certain mountains or islands (Inger, 1954, Inger, 1954, andBrown and Alcala, 1955,1970a). In an addi- 1999; Leviton, 1963; Brown and Alcala, 1970a). tional phylogenetic study, Brown et al. (2000c; see Brown Published fauaal papers are extremely important and Diesmos, in press, for review) conducted a phylogenetic because of their role in educating the international analysis of the flap-legged geckos, genus Luperosaurus (half community about Philippine biodiversity, and be- of which are Philippine endemics). This study showed evi- cause they are an important source of baseline data dence of two monophyletic clades, one with three non-Phil- for bio geo graphers, conservation biolo gists, ecolo- ippine species and the other containing the four Philippine gists, and systematists. Unfortunately, many impor- species plus one species from northern Borneo. The position tant data that have been collected are unavailable in their of the Bornean species, nested well within this second clade, unpublished form (government and non-government suggested a re-invasion of Borneo from a Philippine source organization or private organization reports). (probably the Sulu archipelago) following the initial radia-

Silliman /ournal Yol.42 No. I 2001 ^Sillirnan,Iournal Yol.42 No. I 2001 JU brown, L)iesmos t:l Alcala I he 5tate ol I'fttlrptne Herpetol.og J L tion in the Philippine (Fig. 6b; Brown et al., 2000c; Brown long to a clade that also contains species from Sulawesi, sug- and Diesmos, 2000). gesting a novel Philippines-Sulawesi connection (Evans et McGuire and Kiew (2001; see also McGuire and al., unpublished data) that have not been previously suggested Alcala, 2000) have demonstrated that flying lizards possess a by biogeographic studies of birds or mammals. much greater (10-12lineages) species diversity in the Philip- Phylogenetic analyses ofseveral other groups of pines than previously thought and that the endemic Palawan Philippine frogs are underway (Brown et al., unpublished species is much more closely related to the true oceanic Phil- data; Evans et al., unpublished data) and similar studies ippine radiation than it is to Sunda Shelf species as suggested of selected Philippine lizard genera are also currently by earlier taxonomy (Fig. 7; contra Musters, 1983; Inger, in progress (McGuire, Brown, and Diesmos, unpublished 1983; Ross and Lazell,1991). It is clear from McGuire and data). Results of these studies are preliminary but con- Ki ew' s (2000) analysis that Philippine D r a c o are derived from tinue to suggest that the unique dispersal abilities of three separate invasions of the Philippines from the Sunda amphibians and reptiles, coupled with their finer scale Shelf (Fig.7). patterns of differentiation on montane centers of ende- Recent phylogenetic analyses of Old-world ratsnakes mism, have resulted in biogeographic patterns that are (Helfenberger, 2001) do not satisfactorily resolve the ques- very different from those postulated traditionally for tion of the monophyly of the Philippine supspecies of Elaphe birds and mammals. (:Coelognathus) erythrura (philippina, erythrura, We believe that amphibians and reptiles repre- manillensis, and psephenoura; Leviton, 1979), but suggest sent excellent model systems for elucidating that some Philippine lineages (designated as subspecies by phylogenetic and interspecific phylogeographic patterns Leviton, 1979) may, in fact, be valid species that are not each characteristic of lower relative dispersal abilities. As other's closest relatives. This study suggests that the rela- such, they should provide a powerful set of tools for tionships of the Philippine ratsnakes may be more interesting distinguishing between hypotheses of vicariance from than previously thought, but that further studies, focussing those of dispersal (characteristic of birds and volant specifically on the Philippine radiations, are needed. Recent mammals). Furthermore, future studies of Philippine phylogenetic analyses of crotaline snakes (Kraus et a1.,1996; amphibians and reptiles should provide a wealth of in- Malhotra and Thorpe,1997,2000) have included one or two formation to biogeographers on differing evolutionary speci,esknown from the Philippines. These analyses suggest processes that lead to their rnique hiogeographical pat- the placement of Philippine radiations within larger groups terns. of species but, as of yet, no exhaustive studies of Philippine radiations of snakes have been forthcoming. Ecological studies of Philippine amphibians and reptiles One additional line of study (Emerson and Berrigan, 1993; Emerson, 1996; Emerson et al., 2000) contained sev- Although there have been important ecological con- eral Plrilippine species of fanged frogs, genus Limnonectes. tributions to the literature in the last decade, a review of stud- These studies indicate that the Philippine members of this ies conducted in the past is necessary because so much of genus are not a monophyletic group, but instead, most be- what we know is based on earlier work. It has become clear

Silliman uloumal Yol.42 No. 1 2001 ^9illiman,Iournal Yol.42 No. I 2001 that amphibian and reptile community structure is strongly erences of many species are available in the publica- influenced by elevational gradients. The general results of tions listed in this section. workers utilizing elevational transect sampling regimes Several important papers of the past 15 years have (Brown and Alcala,196l; Brown et al., 1995b; 1996,2000b; expanded our knowledge of specific habitat preferences. Diesmos, 1998; Ferner et al., 2001) suggest that species di- Alcala and Brown (1987) discussed the habitat preferences versity decreases and endemicity increases with elevation of the unusual Philippine endemic ftog, Barbourula (with a possible mid-elevation species bulge in diversity; busuangensis. Gonzales and Dans (1994) expounded on ar- Brown and Alcala,1967; Diesmos, 1998)- At present we lack boreal habitat preferences of certain lizards and amphibians the kind of fine scale information on elevational gradients on Mt. Makiling (see also Das and Charles, 1994; see also that has been provided for mammals (e.g., Heaney and Rickart, Torres, 1955), and Gaulke (1995b) reported on the unusual 1990; Heaney et al., I99l; Rickart et a1.,. 1991; but see utilization of arboreal habitats by typhlopids (see also Taylor Diesmos, 1998), and we have no detailed information (other 1922e). Diesmos (199S) gave detailed descriptions of frog than percent endemism) for community structure variation microhabitat preferences on Mt. Makiling and Mt. Banahao' along elevational gradients on land-bridge versus oceanic is- S. Luzon, and Brown et al. (1996, 2000b) have presented lands. Such studies are greatly needed. habitat information on populations in the Zartrbales and Si- erra Madre mountains. Recent survey work by Ferner et al. Habitats.The first sources of habitat preferences of Phil- (2001) and Gaulke (2001a, 2001b, 2001c) includes signifi- ippine amphibians and reptiles have been the descrip- cant new information on the habitat preferences of several tions of the habitats in which species were collected by poorly known species from Panay Island. A recent investiga- taxonomists. Most of the taxonomic works of various tion into cave habitats (C. Dolino. unpublished data) should workers (see Literature Cited; papers by Taylor, Brown, provide interesting new information on subterranean species' Alcala, Rabor, Inger, Leviton, Diesmos, Brown, habitatpreferences (see also Brown and Alcala, 2000). Brown McGuire, Gaulke, Ferner, and collaborators) mention and Diesmos (2000) discuss the paucity of information on specific microhabitats from which specimens were col- canopy habitats in the Philippines (see also Lowman, and lected. From these works we can discern that important Nadkarni, 1995) and the lack of knowledge regarding the microhabitats for amphibians and reptiles collected in microhabitat preferences of geckos of the genera original forests include streamside microhabitats (on and Luperosaurus, Pseudogekko, and Ptychozoon (see also Brown under rocks, overhanging vegetation, debris on the et al., 1997, 200Udiuffenberg and Auffenberg ( 1 9 8 8 ) have banks, etc.), trees (on trunks, in branches, under bark, provided detailed habitat descriptions for 11 sympatric spe- in canopies), epiphytes (aerial ferns, pandans, orchids, cies of southern Luzon scincids, and Auffenberg (1988)' 1 provided moss mats, suspended debris), litter and humus layers, Gaulke ( 1 9 89a, 1992b),and Bennett (1999 a, 999b) upland moss accumulations, etc. A comprehensive syn- some information on varanid lizard habitat preferences. thesis of all that is known about habitat preferences More detailed descriptions of species partitioning in would be very useful, but to date such a reference is heterogeneous habitats and elevational gradients are avail- still lacking. Fortunately, data on the microhabitat pref- able in Alcala (1 9 67, 1 98 0), Custodio ( 1 9 86), Auffenberg and

Silliman,Iournal Yol.42 No. I 2001 Silliman "/ournal Yol.42 No. I 2001 34 Brown,l)iesmos ti Al.cat-a

Auffenberg (1988), Brown et al. (1995b, 1996), Diesmos, boreal foam nests coupled with aquatic development at later 1998; Hampson (1999b). and Ledesma (1999). Additionally, larval stages (Rhacophorus and Polypedate.s; Alcala, 1962: Smith (i993a, 1993b), Alcala and Brown (1998), Bennett Alcala and Brown,1982,1994; Brown et al., 1997a). Most (1999a, 1999b), Hampson (1999a, 1999b, 2001), Ledesma non-platymantine ranids, bufonids, microhylids, megophryids, (1999), and Gaulke (1992b;1994a,1995b, 1996,1999), all and caecilians rely entirely on indirect aquatic development contain other incidental habitat preference details for species (Taylor, 1920a; Inger, 195.1; Alcala and Brown, 1956; Alcala involved. Brown et al. (2000a) utilized microhabitat prefer- and Alcala, 1980; Brown and Alcala, 1982b) while some ence differences to facilitate the recognition of a new species ranids undergo terrestrial development in nests near or arvay offrog from the Siena Madre mountain range (Rana tipanan). from water (Inger, 1954; Alcala, 1962; Brown and Alcala, 1982b; Inger et a1., 1986; see also Brown and Iskandar, 2000). Reproduction and development.There has virtr:ally been no Finally, some life histories in the Philippines still cornpletely progress in the study of developmental biology of Philippine unknown (i.e., Barbourula busuangensis; family species in the past 10 years and nearly all of what we know Bombinatoridae; Taylor and Noble, 1924; Myers, 1943; comes from the studies of earlier workers, most notably A. Brown and Alcala, 1982b; Alcala and Brorvn, 1987; Ubaldo, Alcala, in collaboration with Brown (Alcala and Rabor, 1957; 1999; Diesmos, Infante, Gee, and Brown, unpublished ob- Alcala, 1962; Alcala and Brown, 1955, 1956, 1982; Brown- servations) provide opportunities for exciting future studies. and Alcala 1982b; see also Brown and Reyes, 1956). Given Auffenberg and Auffenberg (1989) provided a detailed the absence of recent studies directed at development and re- descriptive study of reproductive patterns in l1 sympatric production, we are left with an attempt to piece together what skink species from the Caramoan peninsulaof southem Luzon. is known from these earlier studies, combined with an effort Their study described a striking level of diversity in clutch to summarize incidental observations from recent works. With composition (egg number and size). parity mode (viviparous the exception of limited developmental data on a few newly- vs. oviparous), and seasonality (rnonth of egg laying) of the described direct developing tiogs of the genus Platymantis reproductive effort in the species studied. It is clear from this (Brown et al., 1997a,1997b), there has been almost no ttew study that we have barely scratched the surface of describing information published on developmental tirning, reproduc- and understanding patterns in reproductive bio lo gy of Phi I ip- tive effort, clutch size, or other basic life history characteris- pine scincid lizards. It is also quite clear that the spectacular tics since the time of Brown and Alcala's (1982b) review. 4lygfstly of reproductive patterns in Philippine scincids pro- For information on particular species, readers are referred to vides unparalleled opportunities for future research. this work. In general, however, we can state that a high de- There is no comprehensive review of Philippine rep- gree of life history variation is exhibited by Philippine tile reproductive modes available, but sorne information on Amphibia. For example, ranid frogs of the genus Platymantis seasonality and reproductive effort can be found in the pa- all exhibit reliance on direct development (Alcala and Brown, pers ofAlcala(1962;1967), Alcala and Brorvn (1967), Brown 1955b, 1982; Alcala, 1962), while some groups (e.9., and Alcala (1970c,1982b), Auffenberg (1988), Auffenberg rhacophorids) possess a variety of reproductive tactics, from and Auffenberg (1988, 1989), and Gaulke (1989a, 7992a, direct development (all Philautus) to the construction of ar- 1992b). Additionally, it would be very useful to compile a

Silliman "Iournal YoI.42 No. I 2001 Silliman.Ioumal Vol. 42 No. I 2001 I IIL .\LULC UJ T ILLLLPLILC I ICI P(LULVEJ J ' reference for reproductive timing, clutch size, and incubation intact virgin forest to beach side habitats), there exists a wide period for Philippine snakes and lizards. These areas are fer- range of habitats, none of which was utilized by all species tile grounds for future research. considered. In fact, physically similar and dissimilar species pairs (Brachymeles samarensis-9. boulengeri, Mabuya Population biologt. Population studies involving Philippine multicarinata-M. multifasciata, Dasia grisia-Lipinia amphibians and reptiles have been traditionally limited pulchella) occupying sirnilar habitats showed evidence of (Alcala, 1955, 1967 , 1970; Alcala and Brown , 1967; Brown ecological replacement. Finaliy, Auffenberg and Auffenberg and Alcala, 7961, 1963c, 1970c). The most in-depth focal (1988) showed no evidence of prey selection or food as a study of a single Philippine species of reptiles is the work of limiting resource. They did show strong evidence of niche Auffenberg (1988) on gray's monitor lizard, Varanus variation based on habitat preferences (variation in diet com- olivaceus,published just over a decade ago. Auffenberg ( 1988) position as a function of the available prey in different habi- provided information on reproduction, life history trait varia- tats), prey item shifts on populations inhabiting both forested tion, behavior, population size and densities, age structure, and open habitats, and temporal variation in diet brought about natural longevity, and diet of V. olivaceus. Since that time, by natural seasonality. Gaulke (1989a, 199Ia,1992a,1992b) has provided some of Recent studies include the investigation into lizard the same data for selected other subspecies of Varanus communities on Polillo Island by Ledesma (1999) and stud- salvator, and Bennett, (7999a,1999b) has supplernented our ies of frog communities by Hampson (1999b,2001). These knowledge of diet, movement pattems, and parasite loads on studies demonstrated that diversity is highest in forested habi- Polillo island populations of Z s. marmoratus and V. olivaceus. tats, or in boundary areas where forest and perianthropic/ag- There are no recent studies on the population biology of Phil- ricultural commensuals coexist. One of these studies demon- ippine amphibians save for Afuang's (1994) study on the in- strated clearly that trog species density and richness increases troduced species Bufo marinus. with increasing distance into the forest away from agricul- ture (Hampson, 1999b, 2001). Community ecologt. There have been only a few studies of amphibian and reptile communities in the past (Brown and Behavior.There have been virrually no behavioral studies in Alcala, 1961, 1963c; Custodio, 1986; Diesrnos, i998; Brown the history of Philippine herpetology, despite the enormous 9!!J296, 2000b; Ferner et al., 2001). Auffenberg elg potential for research offered by Philippine populations of Auffenberg (1988) provided a detailed description of a com- amphibians and reptiles. There have been sir-,.'iiicant munity of 11 species of sympatric scincids on the Caramoan behavioral observations of selected species, mostly having to Peninsula of S. Luzon. Their analysis showed that scincid do with antipredatory behavior and habitat preferences (Brown species diversity is positively associated with density of veg- and Alcala,196I,1978; Brown et al., 2000a, 2000b), repro- etation and structural complexity and that, among habitats, ductivebehavior (Alcala et al., 1987; Auffenbe4g, 1988;Gaulkg intact original forest was the habitat that supported the high- l99la, I992b; Auffenberg, 1988), diets @eyes , 1957 , 1968), or est species diversity. In natural habitat gradients, such as the even spacingpattems and pattems ofmovement (Auffenberg, I 988; study areautilized by Auffenberg and Auffenberg (1988; from Auffenberg and Auffenberg, 1988; Bennett, 1999a, 1999b)'

^Silliman./ournal Yol.42 No. 1 2001 Silliman../ournal Yol.42 No. I 2001 Snte of PhiLipine Herpetolag; 39 38 Brorun, Die.smos E Alcakt TIw

Recently. there have been an increasing number of 1992; Primak and Lovejoy, 1995). Although logging in papers containing information on communication in Philip- the Philippines has significantly slowed, it is clear that pine frogs (e.g., Alcala et al., 1986; Brzoska et al., 1986; this trend is due primarily to the absence of significant (Heaney Brown et al., 1997b, 1997 c,1999a, 1999b; Hampson, 1999b), stands of Philippine timber left to cut et al.. and one in-depth study of the evolution of diversity of 1999) rather than as a result of government grassroots to behavioral mate-recognition signals in the genus Platymantis wildlife protection initiatives or government efforts currently is underway (Brown et al., unpublished data). sustainably manage resources (Kummer, 1992; Sajise et al., 1996). Conservation: a review of what we know and suspect The last ten years have seen an increase in designa- tion of protected areas and in public awareness of the need to It is abundantly clear that amphibian and reptile preserve the habitats of endangered Philippine amphibians populations in the Philippines are imperiled due to massive and reptiles (Brown and Alcala, 1986; de Celis, 1995; Sajise loss of their forested habitats (Brown and Alcala, 1986, 1994; et al., 1996;DENR and LINEP, I 997; DENR and PALF, I 998: Auffenberg, 1988; Diesmos, 1998; Gaulke, 1989b, 1992b, Heaney and Regalado. 1998; ECPF, 1998; Gaulke, 1998: 1998; Hampson, 1999b; Brown et al., 2000b; Ferner et al., Hicks, 2000; Tan, 2000). These advances in the potential fbr habitatprotection are most encouraging (reviews: Heaney and 2001 ; Heaney and Regalado, I 998; Heaney et al., 1 999). Other anthropogenic factors include the indirect effects of industry Regalado; Heaney and Mittermeier, 1997; Heaney et a1., and population growth, subsistence farming and habitat modi- lee9). fication, and the direct causes of population declines due to years has bcen over-hunting, and exploitation of populations for food and Conservation status of species. In recent there trade (Seale, 1917;Taylor,1920b; Domantay, 1953; Punay, a first genuine attempt to arrive at a consensus concerning 1975; Ross, 1982; Bacolod, 7984, 1990; de Celis, 1995, the conservation status of amphibians and reptiles in the Phil- Gaulke, 1998). Still, despite all other known causes of de- ippines (Magbanua, I 99 I ; Alcala and Custodio, I 995; Afuang clines, we must accept that the removal of original forests or and Gonzales, 1997; Gonzales etal.,l997;CI, FFI, and IUCN- past, other forms of habitat loss remains the most pervasive cause SSC, 1999; Gaulke, 1998; Banks, 1999). In the interna- tional attention, concern, and attempts at regulation in the of population decline in all forms q{ lgqestrial Philippine tur- wildlife (Brown and Alcala, 1986; Whitrnore, 1984; Whitmore form of CITES-orIUCN listings were limited to marine a, and Sayer, i992; Primack and Lovejoy, 1995; Heaney and tles (genera Er etmochelys, Lepidochelys, Che lonia, Crtrett lizards (ge- Mittermeier,1997; Heaney and Regalado, 1998; Heaney et andDermochelys; see also de Celis, 1995), sailfin al., 1999). There can be no doubt that a significant percent- nus Hydrosaurus), a few freshwater turtles (genera Heosentys, age of habitat loss is related to government-sanctioned com- Pelochelys), crocodiles (Crocodylus porosus and C' mercial industries (Heaney and Mittermeier, 1997; Heaney mindorensis; Ross, 1982; Trono ,1997; Ross and Alcala, 1993; pythons and Regalado, 1998; Heaney et a1.,1999). Philippine forests Palma, 1993; Ortega et al., 1993; Regioniel, 1995), continue to be felled at an alarming rate (Bawa et al., 1990; (Py thon r e ti cu I atw), I arge water snakes (e. g., gener a C er b e r us . Whitmore 1990; Collins et al., l99|.Whitmore and Sayer, Acrochordus, Laticauda, Hydrophis and Lapemis), a f-er,'''

,Silliman uloumal Yol.42 No. 1 2001 Silliman..Iournal Yol.42 No. 1 2001 40 Browrr, Die.smos E Ahaln Th.e State oi Philipine Herpetolngl 4l

terrestrial snakes (e.g., genera Naja, Elaphe, Stegonotus, Explo itatio n and con sumption oJ' a mphibians and reptiles. Zoacys; Alcala, 1986; Ross et al., 1987), and monitor lizards There exists only a handful of studies documenting the ex- (Varanus; Gaulke, 1998)-those species presumably at risk ploitation of amphibian and reptile populations (as food due to an aggressive SE Asian leather trade (reviews: unpub- sources, and for the leather and pet trades) in the Philippines. lished Sagip Wildlife Program list; Alcala, 1986; Gonzales In general there are a few published reports that mention the et al., 1997; Erdelen, 1998; Gaulke, 1998; van Dijk et al., use of amphibians and reptiles as food sources by indigenous 2000). More recently, Alcala and Custodio ( I 995) and Afuang groups (Villamor, 1990; Luxmoore and Groombridge, 1989; and Gonzales ( I 997; see also Banks, 1995 , 1999) have begun see also Gaulke, 1992b,1998). We know that amphibians (rice an effort to address the conservation status ofother, less no- field frogs of the genus Rana and fanged river frogs of the ticeable species such as frogs (Afuang and Gonzales, 1997; genus Limnonecfes), reptiles (lizards of the genera CI, FFI, and IUCN-SSC, ;?13; Banks.l999;review: Hilton- Hydrosaurus and Yaranus), and snakes, (i.e., genus h)thon) Taylor, 2000). In contrast to many species status initiatives of form an important part of the diet of many indigenous cul- the past that have argued for increased protection due to over- tures in the Philippines (Lopez,l976; Kikuchi, 1984; Griffin exploitation by humans, more recent projects (Alcala and and Estioko-Griffin, 1985; Schult, 1991; review: Gaulke, Custodio, 1995; Gonzales et al., 1997;' Banks, 1999) show 19S9b). Road-side hawkers offering pythons and monitor liz- that the majority of the newly listed species are considered ards for sale are a common sight throughout the country (ex- threatened primarily by habitat loss, or are vulnerable as a cept in predominantly Muslim areas; pers. obs.\. consequence of limited geographical distributions. However, many of the desired data (species identi- The 1997 Wildlife Conservation Society of the Phil- ties, numbers of individuals harvested, seasonality of harvest, ippines Philippine Red Data Book (WCSP, 1997) represented locations of prirnary harvests, percentage of the harvests that the first attempt to arrive at a consensus as to the conserva- are subadults, sex of specimens harvested) are still lacking. tion status of Philippine amphibians and reptiles. Two am- We are in drastic need of these types of data in order to irn- phibians and l0 reptiles considered globally threatened in the plement informed management decisions. Although leather Philinnines were included. Later, following the launching of and pet trade harvest and export were completely banned in the "Global Arnphibian Carr-aign" (CI, FFI. and IUCN-SSC, 1994, the industry continues to thrive (Bacolod, 1984; 1990; (1999) 1999), Banks included an additional 32 species of Gaulke, 1989b, 1998) and is possibly growing (F. Yuwono, Philippine amphibians in the 2000 Red List of Threatened pers. comm.). Hides of Philippine reptiles continue to appear Species. A new, comprehensive re-assessment of amphibian in overseas markets at the same time that rare and protected species' conservation status will soon be forthcoming Philippine species are now increasingly advertised for sale at (Diesmos et al.. unpublished). We hope these efforts will re- exorbitant prices on the intemet (Brown, pers. obs.) as curi- sult in increased protection of vulnerable populations, in- osities and "captive biological specimens" (: pets), report- creased public awareness (Afuang et al.,2002), the designa- edly, but doubtfully, bred in captivity in an attempt to "legal- tion of conservation priorities based on data (not politics), ize" the selling of protected rvildlife. We do know that un- and increased use of conservation resources towards the study regulated exploitative harvests of sea snakes for skins have and protection of potentially threatened species. devastated rookeries in the Visayan sea (Bacolod, 1984; 1990).

Silliman "fournal Vol. 42 No. 1 2001 .Silliman,Iournal Yol.42 No. I 2001 The State of Philipine Herpetology 43 42 Bto*n, Diesmos E Abahr

future yields and their own and that at present there are no specit-lc laws in place to pro- over-harvesting in order to insure of reptil e harvests would tect sea snakes from leather trade overexploitation (Gaulke, livelihood. Finally, legal monitoring data for policy makers 1989b). The next decade will be a critical period in which the provide a great many badly-needed on yields, size of challenges of gaining information on these uses of amphib- and wildlife biologists. with information and harvest locations (e.g., ians and reptiles must be addressed in a meaningful fashion. harvests, percentages ofeach sex, sound recommen- Some of the countries surrounding the Philippines Shine et al., 1998), informed, biologically decisions could be made to insure have made efforts to monitor, regulate, and sustainably man- dations, and management economically important species age reptile harvests (Erdelen, 1998; van Dijk et al., 2000)' the continued survival of Gaulke, 1998)' In the absence and it is now time to begin a dialogue on the Philippines' (Yuwono, 1998;Melisch, 1998; ignorance and forced to pro- own response to these growing industries. Gaulke (1998) rec- of such data, we are left with unregulated black market in ommended the implementation of regionally-oriented wild- ceed from guesswork, while an continues to thrive' life management plans which include protection of certain Philippines amphibians and reptiles areas, but with legal trapping based on quotas and the princi- the threat they pose Fte' ples of sustainable yield in others. This proposal is worthy of Introd.uction of exotic species and ' (1998: unpublished data; see consideration because of the manner in which it may benefit cent survey work by Diesmos augmented data on Asian and both the animals and the local communities. In general, regu- also Diesmos, 2000, 2001) has the Philippines' We now lated, sustainable harvest of protected species is more desir- American species introduced into Sunda Shelf species Rana able than unfettered, unregulated, unmonitored rarnpant ille- know that in addition to the Alcala, 1986), middle gal exploitation (Webb and Vardon, 1998; Shine et al., 1998; erythraea (Brown and Alcala, 1970c; (Bufo marinus; Alcala, 1986; Afuang, Erdelen, 1998; Webb and Vardon, 1998). We expect that some American cane toads gs (Hop lobatr achus rugulo sus ; Varanus, Acrochordus, Hydrophk, Uticauda, Naia and Py- lgg 4), Taiwanese bullfro and Brown, 1998), and American bull- thon populations can be harvested at sustainable levels once Diesmos, 1998; Alcala Inovejas and Vergara, 1985), have data are available to indicate the appropriate levels and har- frogs (Rana catesbiana; the Philippines. All of vest times. Data needed include the number of individuals established breeding populations in (Diesmos and Brown that can be sustainably harvested from a population, when these species have rapidly spread 'pers' original introductions. The rapid the appropriate (non-breeding) harvest season shQuld oqcur, obs.) from the points of their voracious dietary habits, and invasive abili- and which populations may be sustainably culled versus which generation time, that they represent seri- must be allowed to recover unmolested. iies ofthe latter three species suggest and habitats they currently Illegal collectors view black market trade as a non- ous threats to the communities syntopic pirpulations of Philippine endemics renewable resource that is best exploited as quickly as possi- inhabit and that soonbe seriously threatenedby these introductions. Basic ble in order to accrue as much income as possible before their may (Heyer et al., 1994) on the spread of illegal activities are exposed. In contrast, legally-registered documentary studies and their behavioral interactions with traders and leather merchants who invest in the monitoring these non-native species are badly needed to document and, hope- oftheir resources tend to protect and guard their sources (see Philippine species of potentially catastrophic invasions' papers in Erdelen, 1998, e.g., Yuwono, 1998) and prevent fully, stem the spread

Yol.42 No. 2001 Silliman,Iournal Yol.42 No. I 2001 ^sillirnan,Ioumal I Thz State of Philipine Herpetology 45 44 Bro*n, Diesmos €d Alcaln

ofbiological resources now also Legal issues, restrictions, and research permits late commercial exploitation apply to biodiversity researchers and field biologists' must also collect and preserve Gaulke (1998) reviewed the laws (or absence thereof) Although biologists part of biodiversity studies, they do governing the exploitation and harvest of monitor lizards, biological specimens as these preserved animals and plants for personal or pythons, sea snakes, and file snakes in the Philippines. The not use gain but, instead, deposit them in internationally passage of Executive Order 247 of the Ramos adrninistration commercial such as the National Museum of the (la Vifra et al., 1997) and the recent Wildlife Bill under the accredited institutions (Simmons, 1987;Reynolds et il., 19941. Resetar Macapagal-Arroyo administration are both new atternpts to Philippines become part of the public record protect Philippine wildlife and natural resources, including and Voris, lgg7)where they heritage of the nation rather than contribute to reptiles and amphibians. These efforts are generally encour- and natural enterprises. The philosophical, ethical, and aging in thai ircy demonstrate an increased concern for the money-making the activities of non-profit, sci- welfare of Philippine wildlife. Unfortunately, the co-occur- practical diftbrences between and for-profi t, commercial bioprospectors ring legal restrictions on the activities ofresearch scientists entifi c biologists the scope of this paper, but numerous obvious and wildlife biologists have seriously crippled biodiversity are beyond are imrnediately apparent. The need for regula- research. distinctions that also distinguishes between the activities At present we see the absence of a clear cut distinc- tory legislation bioprospectors and research biologists should tion between academic/research and commercially-oriented of commercial Finally, although we are aware that it activities (La Viffa et al., 1997) as a policy in need of revi- be equally apparent. was not intended as such, the current implementation of EO sion. Without such a distinction, EO 247 will continue to crip- amounts to a policy of economic discrimination against ple biodiversity studies, despite its good intentions. One nega- 247 students and junior scientists. This is because the tive impact of the passage of EO 247 has been the tnanner in university endless lists of legal requirements make it pro- which it has contributed to an incorrect public perception of seemingly expensive and nearly impossible foruniversity stu- biologists as somehow akin to commercial exploiters of the hibitively working with modest budgets to obtain environment ("bioprospectors"). Executive Order 247 was dents and biologists research permits. designed to protect wildlife and the Philippine environment legitimate suspect that rnost of the present bureaucratic re- from commercially exploitaf'rveenlerprises such as large scale --W€ stem from the understandable yet un- commercial harvesting of wildlife (i.e., butterfly and orchid strictions on biologists policymakers that the best way to pre- collecting for lucrative overseas markets, unregulated pet trade informed opinion of wildlife is to prevent any killing of animals, harvests, or large-scale collecting ofsnakes, lizards, and tur- serve Philippine name of identifyrng and cataloging the country's tles for shell and leather trades), commercial pharmaceutical even in the is to find fault with these sentiments extraction of potentially valuable plant extracts, ccmmercial biodiversity. It difficult we too disdain the needless killing of animals' How- logging, or any other activity on the part of persons or groups because prevention of responsible faunal collecting ef- who would profit frorn the sale or copyright of Philippine ever, the total forts as part of legitimate biodiversity studies is misdirected. biological resources (La Vifra eta1.,1997). Unfortunately, the there is simply no substitute for vouchered locality data same restrictions that were developed to monitor and regu- First,

2001 Cillimen /nrrrrrol \/n1 4) lrTn 1 ?OOl Silliman../ournal Yol.42 No. I -I'lw HerpetoLogy + I 46 Bro*n, Diesmos E Alraln State ol I'hiLiptne for mapping distributions of species (Reynolds et al',1994). Comparisons lvith neighboring countries Second, there is no evidence to support the notion that re- sponsible scientific collecting has negative impacts on natu- A superhcial look at the herpetological literature from ral populations (Hedges and Thomas, l99l; Goodman and sunounding SE Asian and SW Pacific countries reveals that great Lanyon, 1994; Stuebing, 1998). Finally, the absolute need for trends in Philippine herpetology fit into the context of a past the data generated by biologists' efforts is undeniable. Truly regional increase of knowledge during the half century. effective conservation programs rely heavily on quality mu- Due to the inequality of progress in all regions, wide-scale Neverthe- seum collections (Hawksworth and Mound ,1997; Hedges and comparisons are impossible at the present time. (and hopefully, Thomas, 1991) and the importance of systematic collections less, some valuable comparisons can be made for conservation efforts is immense (Hoagland, 1989; Foster, are heuristic). For example, while estimates of numbers of (from 1982; Nielsen and West, 1994; Savage, 1995; David, 1996; amphibian species in the Philippines have increased 1986; Alcala and Leh, I99 6; Resetar and Vori s, | 997 ; Shaffer et al., 1 99 8 ; Pon- 55 to 105 species; Inger, 1954; Alcala, so too have spe- der et a1., 2001). This is because the baseline data contained Brown, 1998; Brown and Diesmos, inpress), in museum collections form a disproportionately large per- cies estimates increased significantly on the island of Bomeo 1985, centage of material in databasing efforts, conservation prior* (from 92 to 138 recognized species; Inger, 1966;Frost, ity-setting activities, and overall conservation of biological 2000; Duellman, 1993; Inger and Tan,1996a,1996b; Inger, resources (e.g., Conservation International's recent Philippine lg99). In fact sirnilar trends can be seen on the islands of Priority-Setting Workshops-based almost entirely on mu- Java, Sumatra, and Bali (Iskandar, 1998; Frost, 1985; 2000)' seum collection data). Duellman, 1993; Inger, 1999; Iskandar and Colijn, At present, some informal discussions have been ini- Sulawesi (Frost, 1985; Duellman, 1993; Iskandar and Tjan, tiated regarding the establishment of a new Philippine gov- lgg6),New Guinea, and the Solomon-Bisrnark archipelagos emment permitting system that would distinguish between (Frost, 1985, 2000; Duellman,1993; Allison, 1996; Brown, 1 though commercial efforts and academic or university-based research, 1997 ; lnger, I 999 ;Alli son and Kraus, 2 00 )' Similarly, in and we are very hopeful that relief will be forthcoming. How- several comprehensive biodiversity projects are still ever, we must stress that current government policies progress, we are aware that estimates of snake, turtle, and (Welch, 1988; need to be revised so that they promote, facilitate, and iizard diversity have substantially increased encawage responsible research on biodiversity rather Welch et al., 1990; Zhao et al., 1988'20O0; Keng and Tat- than strongly inhibit, restrict, or prevent it. Without such Mong, 1989; Matsui et al., 1989; Cox, 1991; Iverson, 1992; changes, current laws will probably continue to promote Lim and Lim,l99};Zhao andAdler, 1993; Das, 1995' 1996b, local paranoia, eventually causing unproductive rifts 1998; David and Vogel, 1996; Dutta and Manamendra- between the government and non-governilIent, univer- Arachichi, 1996; Inger and Tan, 1996a, 1996b; Inger and sity, local, and scientific communities. The result of such Stuebing, 1989, 199? Chou and Lin, 1997; Manthey and rifts can only be that Philippine environment, Filipino Grossman, 1997; Cox et al., 1998; da Silva, 1998; Chan-ard et biologists, and the biodiversity of this country will con- et al., 1999; Inger,1999; Liat and Das, 1999; McDiarmid tinue to suffer. al, 1999; Ota, 1999; Stuebing and Inger, 1999; Iskandar,

Silliman -Ioumal Yol.42 No. 1 2001 .Silliman.Iournal Yol.42 No. 1 2001 48 Bro*n, Diesmos E Alcala Tlre Srate of P|'ciLipine HerpetoLogl *Y

2000). Although a comprehensive review of all types of stud- knowledge of amphibian diversity in this isolated moun- ies involving amphibians and reptiles throughout Asia and tain range (see comments by Diesmos, 1998). Recent the Pacific is beyond the scope of this paper, our general im- work in the Central Cordillera (Heaney et al., 2000; pression is that the same trends that we have witnessed in the Diesmos, Brown, Gee, unpublished data) should provide Philippines, specifically an explosion in the types of studies an important prelirninary update towards the assessment and dramatic increase in biodiversity and conservation, have of this mountain range's herpetological fauna, but other occurred throughout SE Asia. As such, progress in philip- localities, specifically in the southern portions of the pine herpetology fits into a broader context of the overall Cordillera, are in equally critical need of similar stud- trends seen in SE Asia: dramatic increases in estimated num- ies. bers of species, increased understanding of natural history, Likewise, the mountains of the Bicol Peninsula systematics, biogeography, and ecology coupled with a dras- each deserve intensive survey efforts (see Brown et al., tic need for more information and conservation initiatives. 2002). Outside of Luzon. numerous other areas require basic survey efforts. These include southeastern Future directions: the decade to come Mindoro, all of Samar and Leyte (but see Gaulke, 1994b: Deuzer et al., 1999), high elevation habitats of Mindanao Targets: species, sites, and kinds of studies.In this sec- (but see Rabor and Alcala, 1959; Smith, 1993a, 1993b), tion we attempt to identify substantive gaps or research and numerous smaller islands including (but not lim- topics in need of study in Philippine herpetology. ited to) Masbate (but see Gaulke, and Altenbach, 1994c), In general, there has been more recent taxonomic Sibuyan, Lubang, Burias, Siquijor, Camiguin, Maestro work in amphibians than in reptiles. Accordingly, while de Campo, Semirara, the Batanes and Babuyans, all of we know of numerous undescribed Philippine amphib- Palawan, Busuanga, Coron (But see Gaulke, 1999), and ians, we suspect that far more numerous species of rep- the Sulu archipelago (but see Gaulke, 1993, 1994a, tiles await discovery. There is a great need for compre- 1995a,1996). hensive reviews of Philippine lizards and snakes within Finally, basic population biology, behavioral, the context of modern species concepts. and reproductive biology studies are needed for Additionally, numerous regions of the philippines numerous species believed to be threatened by ac- cry out for faunal surveys. In a recent faunal survey in tivities of hurrrarrs. {t is only through the caret-ul Aurora Memorial Natural Park, Brown et al. (2000b) collection of basic population and demographic data stressed the need for exhaustive herpetological surveys that we will be able to make sound management throughout the Sierra Madre range. Similarly, while recommendations. And it is only through the col- Brown et al. (1996) have provided a preliminary account lection of basic data on the use of amphibians and of herpetological communities in the Zambales, their reptiles by commercial and indigenous harvesters survey was conducted immediately following the erup- that we will be able to assess which populations are tion of Mt. Pinatubo, and so we suggest that further sur- being most heavily exploited. veys are needed, especially if we are to gain an adequate

Silliman,/ournal YoL42 No. 1 2001 Silliman -Iournal Yol.42 No. I 2001 50 Brown, Diesmos I Atcal^q The State of Philipine HerpetoLogy 5l

Publications and survey data. One final lesson from our Field work. As suggested by Crombie's quote at the be- experiences over the past decade that cannot be stressed ginning of this paper, we believe the degree to which too often or too fervently is the need to encourage stu- basic reliable distribution data are lacking and badly dents, government, non-government, and even con- needed cannot be stressed too often. Unfortunately, the tracted workers to publish the results of their studies. public disinterest, financial difficulties, and bureaucratic The amount of critically important unpublished data that obstacles faced by any budding tield research program we are aware of is staggering. If the information con- in herpetology at the present day in the Philippines can tained in non-government organizations' and university be overwhelming. To students tinding thernselves in students' unpublished reports was now available to wild- these or similar situations we wish to offer our encour- life managers, conservation biologists, biodiversity spe- agement and assistance wherever possible. This is be- cialists, and biogeographers, the state of philippine cause a comprehensive, careful, and well-orchestrated herpetology would be markedly different than it is at (and published in a timely fashion) field survey of even present. In truth, unpublished survey data may do more a single forested site makes a major contribution to our harm than good because the tendency is for permitting collective knowledge of Philippine herpetology. Simple authorities to discourage reinvestigations of previously- - "rice and beans" (Crombie, 1992) or "bean-counting" surveyed areas. Thus, unpublished data not only are (A. Malliari, pers. comm.) field exercises can drastically unjustified (why collect data if they will not be put to change the way we view complex topics such as the in- use as part of the public record?), but they actually have fluence of geological processes and marine barriers to a negative impact by barring later workers access to the gene flow on speciation and the composition of faunal same regions (Crornbie, 1992). communities, the effects of elevation on species abun- Similarly, rushed or non-exhaustive, or even dance and distribution patterns, and overall the burgeoningly popular "rapid assessment" sur- zoogeographical relationships of particular islands veys can often do more harm than good. In this in- (Brown et al., 1996;2000b; Diesmos, 1998; Ferner et stance, "a little" is not "better than nothing at all" al., 2001). For all of these data, and the paradigm-alter- if the results are that permitting authorities deny ing conclusions that have been, and continue to be drawn permission to conduct follow up surveys because from them, there is no substitute for reliable distribu- the percepT-ron iS that the work has already been tion data based on specimens deposited is aeeredited completed. No amount of reanalysis of insufficient natural history museums. data will have positive or even illustrative results. We agree with Crombie's recent comment that Integration. We anticipate that the next decade will see "...considerable money and effort are being ex- a genuine effort to integrate recent efforts of taxono- pended on analyzing [herpetological species] dis- mists, systematists, biogeographers, and conservation- tribution information when the data base is so pal- ists. Our review of the iiterature suggests that current try that it scarcely warrants the exercise" (Crombie, herpetology in the Philippines is in a final stage of dis- 1992:594). covery. This descriptive, piece-meal process will no

,Silliman.Iournal YoL 42No. I 2001 .Silliman,Iournal Yol.42 No. 1 2001 nerpetowg1 52 Brown, Diesmos E Alral^a l'fu State of I'fvtLtptne rJ doubt culminate in the availability of an enormous learned through partnerships with local communities. In amount of data available for reviews, syntheses of tax- one sense, local communities are the most important onomy and distribution, large scale biogeographic stud- guardians of the remaining forests; as such, it is in eve- ies, and meta-analyses of ecological studies. Ecologi- iyon"'t best interest that scientists, government officials, cal, behavioral, and population studies will no doubt regional resource managers, and local indigenous peo- contribute to conservation if they can be integrated into plis' organizations work together. The most productive larger synthetic analyses within the context of known research programs of recent history have all been col- history. Recent comprehensive studies of taxonomy, sys- laborative efforts. By combining efforts, Filipinos and tematics, and the numerous factors affecting species dis- non-Filipinos have been able to achieve much more in tributions will no doubt have broad implications for collaboration than could have been possible as part of conservation and management decisions. Integrating separate research programs' We are greatly encouraged these studies and formulating and implementing poli- by the fact that recent collaborative research efforts and cies on the basis of sound biology (instead of politics) conservation prograrls are now being led by Filipino will be a major challenge for the next decade's biolo- biologists. gists, students, and policy makers. The last decatle and the next generation of Philippine Collaboration. It is instructive to note that Philippine herpetologisrs. This last decade has left us with a grow- herpetology has a rich recent history of international col- ing sense of urgency and the ever-increasing need to laborative efforts. In particular, the development of Phil- involve and encourage Filipino students to participate ippine herpetology since the 1950s has relied, at least in the study of their country's amphibians and reptiles' in part, on foreign support. It has been this cooperation In particular, we are encouraged by the recent emergence and partnership ofscientists from several different coun- of numerous women in Philippine herpetology and we tries that has produced the most remarkable discoveries support their interest and involvement in a field of sci- and advances in Philippine herpetology. This tradition ence traditionally dominated by a few male personali- has taken the form of financial support for field research, ties. We are intrigued to imagine who will constitute advice, guidance, encouragement, and facilitation of the next generation of Philippine herpetologists and we aCademic studies abroad. We feel this history proVides wish to encourage all jslerested students to pursue us with an important lesson. Biodiversity studies by both herpetology as a field of study, especially in the field, Filipinos and foreigners should be conducted in collabo- even (and perhaps especially) as represented by the ration with Filipinos at all levels-government, univer- populations in their backyards. It is our hope that the next sity, municipality and the barangay. Our experience has generation of Philippine herpetologists can learn from our shown that it is in the best interest of everyone for re- trials, our accomplishments, and our mistakes' and continue searchers coming to the Philippines to collaborate to work towards new, ever-enlightening conclusions' We hope closely with Philippine scientists and local community students will find inspiration frorn past achievements in the representatives. There is a great deal to be shared and field to rcaltzetheir own power to make significant contribu-

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1990' Table 1. List of amphibian species described since

D is tribution Species F arnily Authoritv lsl. and Gonzales, 1992 C atanduanes Ktloula kokacii lvl rc rohylid ae Ross hit. Hilong-hilong, Mindana': lsl Rhacophoridae Brorryn and Alcala, 1994 P hilaulus Poecilts Peak, Ivltldarrao Isl and Alcala, 1994 Daptten surrufus Rhacophoridae Brown l-7 Phiiautus 199? til t. l',{ adja-as, FanaY lsl Ranid ae Errr'*,n, Bro'cn, and Alcala, P l( ry E a n tis Pa naYe nsis h{ t. lsarcg, Luzon lsi Br0wfl, Br0!vD, Alcaia, and Fro tl" 199'l E PIalymaitis isarog R anid ae 0t td aquiling, Luzrn l::l' tr, Alc ala, and Dre stro s' I 9 9 ? IU t Flatyi2antis mirnulus Ranid ae Brrrw and Alcala' 199? C antaub. Bohol lsl o R alid ae Erowrr, Alcala, Diesnos. P lalvm anlis rabc'ri fl egras Isl I and Alcala' I 9 9? Cr:ertio s de l{ egros, : ae Brlv.,n, Alcala, Diesmo s, a P la i)' il d n tis ne gro se n sis Ralid tD lsl c s, and Alcala' I 9 9? I{ t }il aquilirrg. Lrrzon Ranrd ae Elrown, Alcaia, Diesmo F n a n lis luzo rse nsis s latv Lu:rn lsl Brorn, Alcala, L)iesmis, and Alcala' 199? Lrlt. Banahao, a o Plalyilantts banahao R anid ae ( :-- ailre rrountairrs, Luzorr lsl' y'.lcala, Bro'rn, arrd Diesmos, 1998 Sierra l,{ Jt ['laly$antis PYgmaeus Ranid at N) Banahao, Luzon Is Brn'wtr, and Diesmos, 1998 I\4t naom iae Ranrdae Alcalr, z Pldt)'manlis Lu:ln lsl ong, and Diesmos' 1999 Sierra tt adtE nirluniaillr, adrensis R atiid ire Brrt,rr. Alcala, f Jal;,n anlis si€rratfl lsl' I i C entral C ord iiie ra mo r)rlta&s, Luzorr Diesrrros, 1999 F anid ae Brou,n, Aicals, and l-) P lalYm anlts caEflYanetxsts M a.Jre fiDurlianls, Luzon lsl O ala, and Die smo s, I 9 9 9 Sierra F" anid ae Brort n, .4L P Iaryn dniis taYlori Lri:'-'rr l:l .{lcala, irrd Diesm0s, I tr99 l{ t. B anal,3r, s lts Ranid ae Ero'ri. P lr n a n lis P s e il d 6 d''' r a i, :: ari t]nstohal' Lrrzorr lsl Dit srro s, i 9I9 Irrl ts. Banalial 3ILJ F arrtd at E;rrw rt, Ak 11a, ald .P /r.l.yri ar Jis iil der''lE r?s!l'( Sierra Nl sdfB fllir.llitaills, L':zln isl l,ro'sn. l.i cGuu:, aurl L)ttsmos, 2000 Fl.atld ii!:andfr F. anid ae dg, *,1r M ndlro Isl R otir:l ae rr,:! i-iuttlri.ill, lil trrless r{.i{"r Ig}r -r11iaIes tu revt;'o S r,rrthern l,rtzrn l:j rcr rlLlhd a: [,ii!:rrtr-,s. t.tru,tl, lld Alcal:, !:l ai':ui't Eev sPstii'J L{ Table 2. List of reptilian taxa described since 1989.

Species Farnily Authority D is trib u ti on \/r o\

EU Lep id od a c ty lu s ba lio b u riu s G e kko nid ae Ota and Crombic, 1989 B atan Isl. oi Draco jareckii Agamidac Lazcll, 1992 Batan Isl. t V2 Typhlops castavolus Typ hlo p id ae Wynn and Lcviton, 1993 Inampulugan Isl. U

p J Typhlops collaris Typ hlo id ae Wynn and Leviton, 1993 Mt. Anuling, Luzon Isl. J SJ o Lycodon alcalai C olubridae Ota and Ross, 1994 B atan Isl. a b Lycodon bibonius C olubrid ae Ota and Ross, 1994 Camiguin Isl. tr Lyc o do ch ry ro C 3 n soprate s olubridae Ota and Ross, 1994 Dalupiri Isl. eH Fo Lycodon solivagus C olubrid ae Ota and Ross, 1994 Central C ordillera mountains, Luzon Isl. o :- Ah aetu lla p ra sin a su lue n sis C olubrid ae Gaulke, 1994 Tawitawi island group, Sulu archipclago 5 NJ Sphe no m orphu s kila n gladensis S cin cid ae Brown,1995 Mt. Kitanglad, Murdanao Isl. z Sphe no m orphu s kno llmanae S cin cid ae Brown, Ferner, and Ruedas, 1995 Mt. Isarog, Luzon Isl. 9 Brachymeles minimus S cin cid ac Brown and E. Alcala, 1 99 5 Catanduanes Isl. N) Parvosciacus sisozi . S cin cid ae Ferner, Brown, and Grecr, 1997 Mt. Madja-as, Panay Isl. O Sph e nc m orphu s ta gap dy o S cin cid ae Brown, McGuire, Ferner, and Alcala, 1999 Mt. Maaling-aling, Luzon Isl.

P se udo ra bdio n lalo nuran C o lub rid ae Brown, Leviton, and Sison, 1999 Mt. Madja-as, Panay Isl.

Draco palawanensis Agamidae McGuire and Alcala, 2000 Palawan Isl.

Hologerrhum dermali C olubridac Brown, Leviton, Ferner, and Sison,2001 Mt. Madja-as, Panay Isl.

i Lycodon /austi C olub rid ae Gaulke,2002 NW Panay Isl. I Varanus mabilang Var anid a c Gaulkc and Curio,2001 NW Panay Isl.

Numbers of publications per year and cumulative totals

N UJ A u O) UJ q o o o O o O O O o c) o i 990

cJ r 991 br:\ 1992 !o !o -uO Jo) 1 993 lJ ('3CC r'o 1 994 6c ^\+ OJ A) :-c) r 995 +'C :foo'<' tJ (t r 996 (t+ .7:.;- .^o C) o0) -ol * 1997 o I 998 o c)r{v 0) ? 1 999 2000

2001 Tfu Stctc oi Plt|ipine Flerperolog 59

(Fig. 2) conrpositiorr of the rast decadr"s ritcrature .' Ilrririppine amphibians (A) and reptiles (B). (Fig. 3) The rclationship bctlrcen the cumulativc total number ol' amphibian specics in thc I'hilippines and the year of dcscription. Note the dramatic increase in rate ol'descriptions in the past decade. 'f hc A Amphibians linal point on this linc (indicated with question rnark) is the estinrated number of new spccies arvaiting description (Diesnros, Ilrorvn, and Alcala, unpublishcd data ). Endemic 120 sOecies tf "

Alcara \ ? 2000 rnger\ \ l,nnu o8o \\,-\ o rayro\ a.f+e Euo \ r*d

J O 2o sprcres

OOO oooo $@N t Systematics and biogeography NN@ E8$3 Taxonomy and I species diversity Year of description Reptiles m Conservation B (Fig. a) The relationship between the cumulative total number of rep- tr Ecology year U Other tile species in the Philippines and the of description. For sinr- plicity, only total species counts (endemic + non-endemic) are sholvn, and these are broken down into snakes, lizards, and turtles. Species diversity in crocodilian species is n = 2. F'inal specics counts (indi- cated with question nrarks) are estimated nurrrbers of new spccies awaiting description.

150

,? o I 1oo c) i! 5 Eso: O

Year of description

S'illiman"/oumal \,'o1.42 No. I 2001 Sillirnan Journal Vol, 42 No. 1 2001 OIJ lJrown, l)iesrnos E Atcokt The State af Pl'tiLtpine HerpetoLogl 6l

(Fig. 5) Formation of gcological conrponcntry of the Philip- (Fig. 6) The preferred phylogenetic hypotheses for the Rana pines: (A: the ma.jor Pleistocene aggregate island cornplexes as iig-n"to of Philippine and Bornean stream frogs "ornpl"" delineated try the 120 rrnderwater bathyrnetric contour and Biowrr, 199i; Browtt Guttman, in pressl bold terminal known mid- to late-Pleistoccne (following "od and sea Ievel rccluctions branches indicate Philippine R' signata complex species) Heaney, 1985, 1986). (B: the preferred phylogen"ti" tt.. for the genus Luperosaurzs Brownetal.,2000c;boldterminalbranchesindicatePhilip- Less than 120 m pine species). submarine contour n. sp.(Mindoro) moellendorffi cf signata sP.2 grandocula similis signata erythraea Tablas + Romblon magnus cancrivora Sibuyan Masbate vittigera

P. intermedium P. rhacoPhorus L. iskandari - L. brooksi L. browni L. yasumai joloensis Borneo L. L. cumingi L. palawanensis L. macgregori

Silliman,/ournal Vol. 42 No. I 2001 ,silliman,Iournal Yol.42 No. I 2001 'l 62 Brown, Diesmos E Abala hz Snte of YrLtLtpme nerpetoLogJ eJ

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Bennett, D. 1999b. Notes on varanus salvator nwrmoratus on polillo Brown, R. M., J. W. Ferner, and A. C. Diesmos. t99i. pefinition of the Island, Philippines. Pp.29-32.In: Bennett, D. (Ed.). Wildlife of Philippine parachute gecko, P ty c lw zoo n inte rnledium T ay lor I 9 | 5 Polillo Island, Philippines: Oxford University-Universiry of the (Reptilia; Lacertilia, Gekkonidae): redescription, designation of Philippines ar Los Bafros polillo Final Report. Viper press, a neotype, and comparisons with related species. Herpetologica Glossup, Great Britain. 53:357-373. Boettger, 1893. O. Neue reptilien und barrachier aus West-Java. hol. Brown, R. M., and S. I. Guttman. In press. Systematic evolution in the Anz 16:334-340. Rana signata complex of Philippine and Bornean stream frogs: Boulenger, G. A. 1882. catalogue of the Batrachia Saliente s Ecaudata in reconsideration of Huxley's modification of Wallace's Line at the collection of the British Museum. London: Taylor and Francis. the Oriental-Australian faunal zone interface. Biological Journal t26pp. of the Linnean Society. Boulenger palawan G. A. 1894. On the herpetological fauna of and Brown, R. M. and D. T. Iskandar. 2000. Nest site selection. larval hatch- Balabac. Annals and Magazine of Natural History g0:6_90. ing, and advertisement calls, of Rana arathootti (Amphibia; Boulenger papuan, G. A. 1920. A monograph of the South Asian, Anura; Ranidae) from southwestern Sulawesi (Celebes) Island, Melanesian and Australian frogs of the genus Rana. Records of Indonesia. Journal of Herpetology 34:444-4 13. the Indian Museum 2O:l-126. Brown, R. M., J. A. McGuire, J. W. Ferner, & A. C. Diesmos. 1999a. A Brown, R. M. 1997. Systematic Evolution in the Rana signata Complex new species of diminutive scincid lizard (Squamata; Lygosominae; of Philippine and Bornean Stream Frogs: Huxley's Modification Sphenomorphus) from Luzon Island, Republic of the Philippines. of wallace's Line Reconsidered at the oriental-Australian Faunal Copeia 1999:362-370. Zone Interface. Unpublished M.S. thesis, Miami Universiry, Ox_ Brown, R. M., A. E. Leviton, & R. V. Sison. 1999b. Description of a new ford, Ohio. ii + pp. 74 species of Pseudorabdiore (Serpentes: Colubridae) tiom Panay Is- Brown, R. M. and A. C. Alcala. 2000. Geckos, cave frogs, and small land, Philippines with a revised key to the genus. Asiatic land-bridge islands in the Visayan sea. Haring lbonZ:19-ZZ. Herpetological Research 8:7 - 12. Brown, R. M. and A. C. Diesmos. 2001. Application of lineage_based Brown, R. M., J. A. McGuire, and A. C. Diesmos. 2000a. Status of some species concepts to oceanic island frog populations: the effects Philippines frogs referred to Rana everetti (Anura: Ranidae), de- of differing taxonomic philip_ philosophies on rhe estimation of scription of a new species, and resunection of R. igorota Taylor pine biodiversity. This volume. 1922. H e rpeto lo g ica 56:8 l - I 04. Brown, R. M., and A. C. Diesmos. 2000. The lizard genus Luperosaurusi Brown, R. M., J. A. McGuire, J. W. Ferner, N. Icarangal, and R. S. taxonomy, history, and conservation prospects for some of the Kennedy. 2000b. Amphibians and reptiles of Luzon Island, II: pre- world's rarest lizards. sylvatrop, The Technical Journar of philip- liminary report on the herpetofauna of Aurora Memorial National pine Ecosystems and Natural Resources l0: lO7-124. Park, Phil ippines. Hamadry ad 25 : I 75- I 9-5. Brown, R. M., J. W. Ferner, and L. A. Ruedas. 1995a. A new species of Brown, R. M., J. Supriatna, and H. Ota. 2000c. Discovery of a new spe- (Reptilia; lygqlqryine lizard Lacertilia; Scincidae;_ cies of Luperoscunrs (Squamata; Gekkonidae ) from Sulawesi, with Sphenomorphr.a) from philippines. pro_ ML Isarog, Luzon Island, a phylogenetic analysis of the genus, and comments on the status ceedings of the Biological g_2g. Society of Washington I 0g : I of L. serraticaudus. Copeia 2000: I 9 I -209. Brown, R. M., J. W. Ferner, & R. V. Sison. 1995b. Rediscovery and Brown, R. M., A. E. Leviton, J. W. Ferner, and R. V. Sison.200l. A new redescription of Sphenomorphus beyeri (Reptilia: Lacertilia: species of snake in the genus Hologenhum (Reptilia; Squamata; Scincidae) from philip- the Zambales Mounrains, Luzon Island, Serpentes) from Panay Island, Philippines. Asiatic Herpetological pines. Proceedings _ of the Biological Society of Washington l0g:61 7. Research 9:9-22. Brown, R. M., J. W. p. Ferner, R. V. Sison, C. Gonzales, & R. S. Kennedy. Brown, R. M., R. Fernandez. C. Rivero, R. Buenviaje, and A. Diesmos. I 996' Amphibians and reptires of the Zambares Mountains of Luzon 2002. Mt. Isarog's herpetological wonders. Haring Ibon 3:12-16. Island, Republic philippines. of the Herpetological Natural His_ Brown, W. C. 1997. Biogeography of amphibians in the islands of the tory 4:l-22. southwest Pacific. Proceedings of the California Academy of Sci-

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